Broad-scale patterns of body size in squamate reptiles of Europe and North America

Aim  To document geographical interspecific patterns of body size of European and North American squamate reptile assemblages and explore the relationship between body size patterns and environmental gradients.
Location  North America and western Europe.
Methods  We processed distribution maps for native species of squamate reptiles to document interspecific spatial variation of body size at a grain size of 110 × 110 km. We also examined seven environmental variables linked to four hypotheses possibly influencing body size gradients. We used simple and multiple regression, evaluated using information theory, to identify the set of models best supported by the data.
Results  Europe is characterized by clear latitudinal trends in body size, whereas geographical variation in body size in North America is complex. There is a consistent association of mean body size with measures of ambient energy in both regions, although lizards increase in size northwards whereas snakes show the opposite pattern. Our best models accounted for almost 60% of the variation in body size of lizards and snakes within Europe, but the proportions of variance explained in North America were less than 20%.
Main conclusions  Although body size influences the energy balance of thermoregulating ectotherms, inconsistent biogeographical patterns and contrasting associations with energy in lizards and snakes suggest that no single mechanism can explain variation of reptile body size in the northern temperate zone.     

Spatial and temporal variation in a suite of life-history traits in two species of wolf spider

Abstract.  1. Life-history traits and density were assayed in seven populations of two sympatric species of wolf spider for three consecutive years. The goal of the study was to quantify population dynamics and its relation to spatial and temporal life-history variation.
2. Adult female body size and fecundity varied significantly, among field sites and among years, in both species. Female spiders of both species differed in mean relative reproductive effort among sites, but not among years. The size of offspring was invariable, with no significant differences due to site or year.
3. All populations of both species tended to either decrease or increase in density during a given year and this was tightly correlated with changes in prey consumption rates.
4. Since life-history patterns are determined primarily by selection, it is concluded that size at sexual maturity for females is phenotypically plastic and responds to changes in prey availability. Offspring size however is not plastic and it is likely that other selection forces have determined offspring size. Temporal fluctuations in population size are correlated over a large area relative to dispersal capabilities for these species and conservation efforts for invertebrates must take this into consideration.     

Asynchronous temporal variation among sites in condition of two carabid species

Abstract.  1. It is generally assumed that generalist predatory carabids are food limited, and the degree of food limitation can vary in space and time, but there are only a few studies that investigate whether or not temporal variation in insect condition is synchronised among sites.
2. In this paper temporal and spatial variation in four fitness-related traits – body mass corrected for size, body size, fat content, and fecundity – of two carabid species, Pterostichus cupreus (L.) and Pterostichus melanarius (Ill.), differing in overwintering strategy, lifespan, diet, and habitat breadth were studied.
3. Interactions between temporal and spatial variation for the fitness-related traits were found for both species. Temporal changes in food limitation were not synchronised among sites, and fitness-related traits were not generally higher on some sites than on others.
4. There was an effect of study year on body mass, fat content, and fecundity for both species, suggesting that temporal environmental variation over large areas is more important than the spatial environmental variation within a year.
5. Within a time period, most different fitness-related traits, except body size, were positively correlated to each other among individuals of the same species. The fitness-related traits of females and males of the same species were generally positively correlated. However, despite the fact that the food resources of the two species overlap, there was no correlation in fitness-related traits between the two species.     

Relative effects of climate change, isolation and competition on body-size evolution in the Japanese field mouse, Apodemus argenteus

Aim This paper examines body size variation in both recent and Quaternary populations of the Japanese field mouse Apodemus argenteus in order to assess the relative effects on body size of climate change, isolation and competitive interactions with its congeneric A. speciosus. Both temporal (since the Last Glacial Maximum, LGM) and spatial (over the Japanese archipelago) scales are considered. Location The small field mouse is widespread in Japan, and the specimens examined were collected from 10 localities on islands of widely differing area (from 4 km² to 230,510 km²) and at latitudes ranging from 30.3° N to 45.1° N. Methods The effects of geographical factors such as latitude and island area on the size variation of A. argenteus were investigated, using the lower incisor size. In addition, the size of some specimens from two Quaternary localities was compared with the size of the extant specimens. Evolutionary rates of size change since the LGM were calculated in darwins. Hutchinson size ratios were used to examine the pattern of variation of the size segregation between the two Japanese field mice, A. argenteus and A. speciosus, in relation to time and space. Results There was a negative relationship between size and latitude among living A. argenteus populations. In addition, there was no effect of island area on body size, especially at higher latitudes. At lower latitudes, A. argenteus were larger on smaller islands, although this trend was not statistically significant. Quaternary specimens of A. argenteus were smaller in size than their living representatives. The interspecific size ratio between the two Japanese Apodemus was larger on smaller islands and at higher latitudes, and there has been a decrease in the size ratio between the two Apodemus since the LGM. Lastly, in accordance with the theory of character displacement, the small A. argenteus was larger in allopatry than in sympatry, whereas the large A. speciosus was smaller in allopatry than in sympatry. Main conclusions These results indicate that A. argenteus does not conform to Bergmann's rule or to the island rule. The variation in size for the small Japanese field mouse at both spatial and temporal scales may be related to climate change, with an additional effect of competition with the large field mouse, especially on smaller islands. The size convergence between the two Japanese Apodemus observed over the last 21,000 years may be explained by the diminution of available food resources due to the reduction of land mass areas following the LGM. It may also be the result of an evolution towards an optimal body size; a hypothesis previously proposed to explain the evolution of body size in island mammals. Lastly, the evolutionary rates of body size calculated for A. argenteus since the LGM are typical of rates calculated for other Quaternary mainland mammals, thus suggesting that the evolution in this species was not particularly rapid, as is often thought for island mammals.     

Reproductive traits of stream-dwelling brown trout Salmo trutta in contrasting neighbouring rivers of central Spain

1. Reproductive traits were studied in seven wild populations of resident brown trout in Spain. We examined whether growth, and certain environmental conditions such as water temperature or food abundance, could explain interpopulation variation in the reproductive characters.
2. The results indicated that nearby populations subjected to a wide variation in environmental conditions exhibited a similar reproductive performance.
3. Age distributions and sex ratios were not significantly different among populations. Age ranged from 0+ to 4+ years but the populations were dominated by the 0+ to 2+ groups. In all rivers females matured at age 2+ as opposed to age 1+ and 2+ for males. Mean length at the end of the growth period differed significantly among populations. However, annual growth rate was similar among rivers and was not significantly correlated with either biomass of the benthos or water temperature during the growth season.
4. Reproductive effort, body condition, fecundity and egg size did not correlate significantly with either the abundance of benthic invertebrates or water temperature. Once the effect of body length on both egg size and number was removed, a significant negative correlation was found between these two traits among populations.     

Stress fever magnitude in laboratory-maintained California ground squirrels varies with season

A previous study demonstrated that California ground squirrels (Spermophilus beecheyi) living in the natural environment had, independent of season, a significantly higher mean diurnal body temperature (T(b)) (39.6 degrees C) than either summer (37.5 degrees C) or winter (36.5 degrees C) laboratory maintained animals. Based upon the previous study it has been suggested that California ground squirrels living in the natural environment may have an elevated set-point for body temperature in a manner analogous to a stress fever response. The present study was conducted to determine if season and/or duration of laboratory open-field exposure influenced the magnitude of laboratory open-field stress fever. If stress fever was involved to some extent in the higher body temperature observed in animals from the natural environment, laboratory maintained animals should exhibit a lower magnitude stress fever during the summer months and a higher magnitude stress fever during the winter months. It was hypothesized that laboratory maintained animals would exhibit the same set-point for stress fever T(b) independent of season, and that the duration of open-field exposure would not influence the magnitude of stress fever. Adult California ground squirrels were acclimated to an ambient temperature of 20+/-1.0 degrees C under either LD 14:10 (summer) or LD 10:14 (winter) photoperiod conditions and individuals from both photoperiod conditions were exposed for periods of 2, 4, and 6 h to an open-field arena. An analysis of the data with a two-factor ANOVA demonstrated that season (photoperiod) significantly influenced the magnitude of the stress fever response (1.1+/-0.1 degrees C for summer animals; 2.1+/-0.2 degrees C for winter animals) while there was no significant influence of open-field exposure duration on stress fever magnitude. These results demonstrate that although the set-point for body temperature in unstressed laboratory maintained California ground squirrels varies with season, the set-point for body temperature in open-field stressed animals does not vary with season. These data lend support to the hypothesis that something like stress fever may play some role in the higher body temperature observed in California ground squirrels living in the natural environment.     

Trypanosomes in American Ground Squirrels

SYNOPSIS. California ground squirrels ( Citellus beecheyi ) were found to be infected with trypa-nosomes whose most striking morphological features are an unusually large kinetoplast and a long, tapering posterior end. The total length is 25.2 μ, and body length is 18.0 μ. This "lewisi" -type species is compared with other trypanosomes reported from American ground squirrels, but a specific diagnosis is not made because the life cycle has not been observed and because morphological features of trypanosomes are known to vary widely under differing environmental conditions.     

Contrasting seasonal dynamics in fleas of the Siberian flying squirrel (Pteromys volans) in Finland

Abstract.  1. The seasonal and spatial variation of the adult flea fauna (Siphonaptera) was examined in connection with live-trapping studies of the Siberian flying squirrel ( Pteromys volans ) in three study areas in southern Finland between 1997 and 2005.
2. The numerically dominant flea species of the Siberian flying squirrel were Tarsopsylla octodecimdentata octodecimdentata and Ceratophyllus (Monopsyllus) indages indages (Ceratophyllidae); the latter being a host specialist of the Siberian flying squirrel. Tarsopsylla octodecimdentata , which also commonly occurs on the red squirrel ( Sciurus vulgaris ), infected a large proportion of the host population throughout the winter, whereas C. indages occurred predominantly during the summer and autumn, being practically the only flea species in nests during the flying squirrels' breeding season and on juveniles still inhabiting their natal nest.
3. The use of nest boxes by flying squirrels did not have any positive effects on flea populations, but it may have had a negative impact on T.   octodecimdentata .
4. The potentially vulnerable C.   i.   indages is a predictable, widespread component in the flea fauna of the Siberian flying squirrel at various spatial scales, and it is likely to occur in most of the permanent flying squirrel populations in Finland.     

Genetic signature of recent glaciation on populations of a near-shore marine fish species (Syngnathus leptorhynchus)

Continental glaciation has played a major role in shaping the present-day phylogeography of freshwater and terrestrial species in the Northern Hemisphere. Recent work suggests that coastal glaciation during ice ages may have also had a significant impact on marine species. The bay pipefish, Syngnathus leptorhynchus , is a near-shore Pacific coast fish species with an exceptionally wide latitudinal distribution, ranging from Bahia Santa Maria, Baja California to Prince William Sound, Alaska. Survey data indicate that S. leptorhynchus is experiencing a range expansion at the northern limit of its range, consistent with colonization from southern populations. The present study uses six novel microsatellite markers and mitochondrial DNA (mtDNA) sequence data to study the present-day population genetic structure of four coastal populations of S. leptorhynchus . Deficits in mtDNA and nuclear DNA diversity in northern populations from regions glaciated during the last glacial maximum (LGM) [ c . 18 000 years before present ( bp )] suggest that these populations were effected by glacial events. Direct estimates of population divergence times derived from both isolation and isolation-with-migration models of evolution are also consistent with a postglacial phylogenetic history of populations north of the LGM. Sequence data further indicate that a population at the southern end of the species range has been separated from the three northern populations since long before the last interglacial event ( c . 130 000 years bp ), suggesting that topographical features along the Pacific coast may maintain population separation in regions unimpacted by coastal glaciation.     

Dispersal and recruitment of Tasiagma ciliata (Trichoptera: Tasimiidae) in rainforest streams, south-eastern Australia

1. This study examined genetic variation within and among populations of the caddis fly Tasiagma ciliata (Tasimiidae: Trichoptera) from rainforest streams in south-east Queensland, Australia.
2. Very low levels of genetic differentiation at large spatial scales, between subcatchments and between catchments, indicated that dispersal by the winged adults is widespread. However, significant genetic differentiation at the smallest spatial scale examined, within reaches in a single stream, suggested limited movement by larvae within streams.
3. A patchy distribution of deviations from Hardy–Weinberg equilibrium and differences in patterns among allozyme loci suggested that populations in particular reaches were the result of only a few matings.
4. These results are surprising, given the large numbers of larvae present within a single reach. We suggest that stochastic effects of recruitment may underlie much of the spatial and temporal variation in population numbers in these rainforest streams.     

Bergmann's rule in larval ant lions: testing the starvation resistance hypothesis

Abstract.  1. Body size of the ant lion Myrmeleon immaculatus follows Bergmann's rule – an increase in body size towards higher latitudes. The hypothesis that ant lion body size is larger in the north as an adaptation for starvation resistance was tested.
2. In a laboratory experiment testing starvation resistance, survivorship curves differed among 10 ant lion populations for both a starved and a fed treatment.
3. The average number of months survived by each population was correlated positively with latitude for both treatments. Across both treatments and all populations, large individuals survived longer than small individuals; however individuals from high latitudes had higher survivorship, even after factoring out variation due to initial body size.
4. These results suggest that starvation resistance may be an adaptation for coping with reduced prey availability in high latitudes. Starvation resistance may contribute to latitudinal gradients in body size of ant lions and other ectotherms.     

Ploidy race distributions since the Last Glacial Maximum in the North American desert shrub, Larrea tridentata

• 1 A classic biogeographic pattern is the alignment of diploid, tetraploid and hexaploid races of creosote bush (Larrea tridentata) across the Chihuahuan, Sonoran and Mohave Deserts of western North America. We used statistically robust differences in guard cell size of modern plants and fossil leaves from packrat middens to map current and past distributions of these ploidy races since the Last Glacial Maximum (LGM).
• 2 Glacial/early Holocene (26–10 ¹⁴C kyr bp or thousands of radiocarbon years before present) populations included diploids along the lower Rio Grande of west Texas, 650 km removed from sympatric diploids and tetraploids in the lower Colorado River Basin of south‐eastern California/south‐western Arizona. Diploids migrated slowly from lower Rio Grande refugia with expansion into the northern Chihuahuan Desert sites forestalled until after ~4.0 ¹⁴C kyr bp . Tetraploids expanded from the lower Colorado River Basin into the northern limits of the Sonoran Desert in central Arizona by 6.4 ¹⁴C kyr bp . Hexaploids appeared by 8.5 ¹⁴C kyr bp in the lower Colorado River Basin, reaching their northernmost limits (~37°N) in the Mohave Desert between 5.6 and 3.9 ¹⁴C kyr bp .
• 3 Modern diploid isolates may have resulted from both vicariant and dispersal events. In central Baja California and the lower Colorado River Basin, modern diploids probably originated from relict populations near glacial refugia. Founder events in the middle and late Holocene established diploid outposts on isolated limestone outcrops in areas of central and southern Arizona dominated by tetraploid populations.
• 4 Geographic alignment of the three ploidy races along the modern gradient of increasingly drier and hotter summers is clearly a postglacial phenomenon, but evolution of both higher ploidy races must have happened before the Holocene. The exact timing and mechanism of polyploidy evolution in creosote bush remains a matter of conjecture.

     

Global change and carnivore body size: data are stasis

Aim  Global warming and other anthropogenic changes to the environment affect many aspects of biology and have often been invoked as causing body size changes in vertebrates. Here we examine a diverse set of carnivore populations in search of patterns in body size change that could reflect global warming (in accord with Bergmann's rule).
Location  Global.
Methods  We used > 4400 specimens representing 22 carnivore species in 52 populations collected over the last few decades to examine whether size changed with collection date when geography and sex are accounted for. We then examined several factors related to global warming, body mass, diet, and the attributes of the different datasets, to see whether they affect the standardized slope (β) of the size versus time regression.
Results  Six of 52 populations we examined show a significant effect of year of collection on body size at the 0.05 probability level. The response of size to global warming does not reflect spatial patterns of size variation, nor do diet or body mass affect tendency of populations to change in body size. Size changes are no more pronounced in populations that have been sampled more recently. However, change, where it occurs, is rapid.
Main conclusions  There may be a tendency in the literature to report only cases where recent changes are prevalent. Although in our data only a minority of populations show body size changes, we may see changes accelerating in the future in response to more drastic climatic changes and other anthropogenic changes.     

Hydrological connectivity drives patterns of macroinvertebrate biodiversity in floodplain rivers of the Australian wet /dry tropics

1. Floodplain rivers in Australia's wet/dry tropics are regarded as being among the most ecologically intact and bio-diverse lotic ecosystems in the world, yet there have been relatively few community-based studies of their aquatic fauna.
2. To investigate relationships between hydrological connectivity and biodiversity in the region, macroinvertebrates were collected from sites within two contrasting floodplain rivers, the 'tropical' Gregory River and 'dryland' Flinders River systems, during the dry season and analysed at various spatial scales. A subset of sites was re-sampled in the following dry season to explore temporal variation. The fauna consisted of 124 morphotaxa, dominated by gatherers and the Insecta.
3. As predicted, hydrological connectivity (the lotic or lentic status of waterbodies) had a major influence on macroinvertebrate assemblage composition and diversity, both in space and time. Assemblages from waterbodies with similar connection histories were most alike, and beta-diversity between assemblages was greatest between lotic and lentic waterbodies, tending to increase with increasing spatial separation.
4. At smaller spatial scales, a number of within-waterbody, habitat and water quality characteristics were important for explaining variation (61%) in the taxonomic organization of assemblages, and characteristics associated with primary productivity and habitat diversity were important for explaining variation (45%) in the functional organization of assemblages. However, much of the small-scale environmental variation across the study region appeared to be related to broad-scale variation in hydrological connectivity, which had both direct and indirect effects on macroinvertebrate assemblages.
5. Conservation of the biodiversity in Australia's wet/dry tropics may depend on conserving the natural variation in hydrological connectivity and the unregulated flow of floodplain rivers.     

Development of an homologous radioimmunoassay for secreted prolactin from the California ground squirrel (Spermophilus beecheyi)

A sensitive homologous radioimmunoassay was developed for secreted prolactin from the California ground squirrel (Spermophilus beecheyi). S. beecheyi plasma and pituitary extracts displaced 125I-labeled S. beecheyi prolactin in a parallel manner with S. beecheyi prolactin (sbPRL). Mean minimum sensitivity of the assay was 0.21 ng/ml, and mean intra- and interassay coefficients of variation were 4.1% and 14.5%, respectively. The assay was used to measure basal prolactin concentrations in male and female ground squirrels at various stages of their annual reproductive cycles. Mean concentrations in nonpregnant, nonlactating young females, pregnant females, and lactating females were 1.63, 11.35, and 10.86 ng/ml, respectively. Mean concentrations in nonreproductive and breeding males were 1.50 and 9.81 ng/ml, respectively. Finally, the assay was used to evaluate cross-reactivity between sbPRL and prolactins and growth hormones from other rodent species. Of the tested hormones, only hamster prolactin showed any cross-reactivity with sbPRL (about 0.03%).     

Body size variation of the floodwater mosquito Aedes albifasciatus in Central Argentina

An inverse relationship between larval density and adult body size has been reported for several mosquito species, affecting their survival and vector competence, response to repellents and other factors. Larvae of the floodwater mosquito Aedes (Ochlerotatus) albifasciatus (Macquart) (Diptera: Culicidae) develop quickly in temporary pools, so intraspecific competition (for food or space) might regulate population abundance and affect the size of adult mosquitoes. We investigated the temporal variation of adult female wing-length (an index of body-size) in natural populations of Ae. albifasciatus, using adults collected during each phase of the rainy season. The relationships between adult mosquito abundance, female wing-length, rainfall and temperature were analysed through simple regressions. Skewness of the frequency distribution of wing-lengths showed a strong negative relationship with mean wing-length. The distribution of wing-lengths varied seasonally and was correlated with rainfall 7-15 days previously as the major consequence of breeding site volume. Thus temporal variation of body size in natural populations of Ae. albifasciatus reflected density-dependent changes in the aquatic habitat where immature stages develop, influenced more by rainfall than by temperature or other environmental variables.     

Ecogeographic analysis of morphological and life-history variation in the Italian treefrog

We describe the spatial pattern of variation in body size, body shape, and adult population age structure of the Italian treefrog (Hyla intermedia). By means of a non-parametric test of matrix association (the Partial Mantel Test), we investigate the role of climate and geography in explaining the patterns of phenotypic variation. Body size is the most important source of morphometric variation, both at intra- and inter-populational levels. Body shape, despite its secondary role in explaining variation, still maintains high discriminative power among populations. Age structure differs significantly among populations, since body size correlates positively to age, these differences partly explain geographic variation of body size. The patterns of variation in both age and body size are congruent with climate variation, suggesting causal relationship. However, we find no evidence for adaptive explanations and suggest that climate-induced demographically differential survivorship might be the proximate cause responsible for the associations observed. Unlike body size, body shape varies congruently with the pattern of between-population geographic distances, which we show to be highly correlated with their genetic distances, thus suggesting isolation by distance as the proximate cause of the association.     

Rodent-flea-plague relationships at the higher elevations of San Diego County, California.

Rodent-flea-plague relationships were examined at sites located at higher elevations of San Diego County during 1991-2002. The most frequently sampled rodents were the California ground squirrel, Spermophilus beecheyi, and white-footed mice, Peromyscus spp. Higher seasonal mean dependent variables (prevalences of squirrels and indices of O. montana), higher mean amounts of seasonal rainfall, and lower seasonal mean ambient temperatures were found at sites during 1991--1993 compared with 2000--2002. Levels of plague antibody indicate that squirrels seroconverted at most sites during spring/early summer when exposed to infected fleas when cleaning out burrows following hibernation, when interacting more with other squirrels occurred establishing home ranges, and when dispersing juveniles occupied abandoned burrows containing old nests. The majority of squirrels (35/41) were reported seropositive at Heise County Park (elev. 1,281 m) during 1996, 2001, and 2002. This may reflect fewer fleas biting squirrels more often when squirrels spent more time below-ground in order to avoid higher ambient temperatures. In contrast, most squirrels (12/13) were reported seropositive at two higher sites (elev. 1,584 and 1,641 m) on Palomar Mountain during 1993--1996 when there were higher mean dependent variables and amounts of seasonal rainfall compared with 1999--2002. This increased rainfall may have benefited host and flea populations and thus plague transmission. Seasonal indices were usually reduced after dusting squirrel burrows with 2% diazinon dust. Findings also indicated that S. beecheyi is a primary reservoir for plague at the higher elevations of these and other sites in southern California, and that the deer mouse, Peromyscus maniculatus, and other Peromyscus spp. were not major host reservoirs at the sites sampled.     

Large-scale synchrony and inter-annual variability in roach recruitment in the Rhône River: the relative role of climatic factors and density-dependent processes

1. We report patterns of temporal variation in the recruitment of roach ( Rutilus rutilus ). The data consist of the annual abundance of the first 2 year-classes, 0+ and 1+ fish, at four sites in the Rhône River (France) between 1987 and 1997. Over this 11-year period both 0+ and 1+ fish fluctuated strongly.
2. Cross-correlation indicated high spatial synchrony in 0+ dynamics, although correlations among sites in 1+ dynamics were weaker. No clear pattern was apparent in the relationship between the level of synchrony and distance between pairs of sites.
3. The spatial synchrony in 0+ fish could be attributable to large-scale variations in weather, influencing water temperature. Total body length of 0+ roach was correlated with water temperature (expressed in degree-days over 12 °C), and water temperature was the main factor explaining inter-annual variation in 0+ cohort size. Monthly variation in abiotic factors (measured by standard deviation in water temperature and discharge) did not influence 0+ fluctuations. Correlations with June water temperature suggest that year-class strength was mainly determined by abiotic factors during the first few months of life.
4. The absence of spatial synchrony in 1+ fluctuations suggests little correlation between survival and abiotic conditions during the first year of life, other factors influencing survival.
5. Survival in the first year was density-dependent. Intraspecific competition within the 0+ cohort could thus influence the fluctuations in recruitment to older age-classes.
6. The implications of age- or stage-dependent synchrony in temporal variation for species with complex life histories are discussed. Studying spatial synchrony for the different life history stages could enhance our understanding of the population dynamics of spatially structured species.     

Glacial refugia for summer-green trees in Europe and south-west Asia as proposed by ECHAM3 time-slice atmospheric model simulations

Aim  To generate maps of potential refugia for summer-green trees during the Last Glacial Maximum (LGM).
Locations  Southern Europe and south-western Asia.
Methods  Time-slice simulations of the atmospheric climate with the ECHAM3 model are used for the LGM. Limiting factors beyond which cool and warm groups of deciduous trees cannot grow (such as temperature in growing degree days, minimum monthly temperature and precipitation in summer) are chosen. A limited validation by fossil pollen and charcoal records from LGM sites was done.
Results  Two sets of maps extending from Europe to the Caspian region for cool and warm summer-green trees are presented. Three criteria are combined using contour lines to indicate confidence levels. Small areas within the three southern peninsulas of Europe (Spain, Italy and Greece) are highlighted as possible refugia for summer-green trees. Further, areas that have remained poorly known are now proposed as refugia, including the Sakarya–Kerempe region in northern Turkey, the east coast of the Black Sea and the area south of the Caspian Sea.
Main conclusions  The maps produced in this study could be used to facilitate better long-term management for the protection of European and south-western Asian biodiversity.